CHUCOA ILICIFOLIA, A SPINY ONOSERIS (ASTERACEAE, MUTISIOIDEAE: ONOSERIDEAE)

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1 Phytologia (December 2009) 91(3) 537 CHUCOA ILICIFOLIA, A SPINY ONOSERIS (ASTERACEAE, MUTISIOIDEAE: ONOSERIDEAE) Jose L. Panero Section of Integrative Biology, 1 University Station, C0930, The University of Texas, Austin, TX 78712, USA panero@mail.utexas.edu ABSTRACT Molecular phylogenetic studies of several members of Mutisioideae show that the type species of the genus Chucoa of northern Peru is a member of the genus Onoseris. The combination Onoseris ilicifolia is proposed. The disposition of the second species of Chucoa awaits molecular studies. Phytologia 91(3): (December, 2009). KEY WORDS: Chucoa, Onoseris, Onoserideae, Mutisioideae, Andes, Peru. Mutisioideae, a taxon first circumscribed based on molecular phylogenetic analysis (Panero and Funk 2008), includes approximately 44 genera and 630 species. The subfamily contains three main lineages recognized at the tribal level: Mutisieae, Nassauvieae and Onoserideae. The monophyly of Mutisioideae has not been confirmed, as a few genera that share morphological features found in this taxon have not yet been included in molecular analyses, and hence our understanding of their relationships remains tentative. Among these unsampled taxa is the genus Chucoa, a seldom-collected species endemic to the Santiago de Chuco region of La Libertad Department in northern Peru. Chucoa was described by Cabrera (1955), who allied the genus to Gochnatia and Stifftia. The same year, Ferreyra named the genus Weberbaueriella to accommodate an unusual specimen of Asteraceae originally identified as an Onoseris. Ferreyra (1955) allied his new genus Weberbaueriella to Gochnatia, Onoseris and Chuquiraga. Chucoa and Weberbaueriella were published in the same year but Ferreyra (1980) acknowledged the priority of the genus

2 538 Phytologia (December 2009) 91(3) Chucoa by placing Weberbaueriella in its synonymy. Both Ferreyra and Cabrera speculated that Chucoa is closely related to Gochnatia. However, morphological studies of Gochnatia and related genera suggested that Chucoa, because of its pubescent styles and acute anther appendages, is not closely related to Gochnatia (Freire et al., 2002). Katinas et al. (2008, Fig. 2) placed Chucoa in an informal group of 22 genera within their concept of tribe Mutusieae, which includes all members of Mutisieae and Onoserideae, sensu Panero and Funk (2008), but did not speculate further as to the sister taxon. Included in this group, because they share a similar style and anther morphology, were also the genera Catamixis, Oldenburgia, Hyaloseris and Dinoseris. However, these four genera have been shown to belong to other lineages of Asteraceae (Panero and Funk, 2008; Panero, 2008). Given these multiple hypotheses of relationship based on morphological studies, we decided to include Chucoa ilicifolia in a phylogenetic analysis of Mutisioideae using chloroplast DNA so as to provide molecular evidence bearing upon the phylogenetic relationship of Chucoa. MATERIALS AND METHODS DNA was extracted from 0.25g dried leaf material of Chucoa, Onoseris onoseroides and Urmenetea using Qiagen's DNeasy Plant Mini Kit following the manufacturer's protocol for dried leaf material. Efficacy of the DNA extract was tested empirically using standard polymerase chain reaction (PCR) protocols and primers detailed in Panero and Crozier (2003) and Panero and Funk (2008) for the following chloroplast loci: matk, ndhd, ndhf, rbcl and rpob. PCR reactions were screened using agarose gel electrophoresis. Reactions with visible results were cleaned using QIAquick PCR purification columns and 4 microliters of each used as template DNA in cycle sequencing reactions following the protocols of the ABI Big Dye Terminator 3.1 Cycle Sequencing Kit. Cleaning of the sequencing reactions using the Millipore MultiScreen 96-Well Filtration Plate and sequencing was performed by the University of Texas ICMB Core DNA Facility on an ABI 3730 DNA analyzer. Raw sequence data were proofread using Sequencher 4.8 (Gene Codes Corporation). Chucoa, Onoseris onoseroides and Urmenetea sequences were aligned

3 Phytologia (December 2009) 91(3) 539 by eye with a subset of 19 taxa of Mutisioideae of the 108-taxon matk, ndhd, ndhf, rbcl and rpob alignments used in previous phylogenetic studies (Panero and Funk, 2008). Phylogenetic trees were constructed for the 22-taxon matk, ndhd, ndhf, rbcl and rpob data sets in combination using the maximum parsimony criterion implemented in PAUP*4b10. Branch support was assessed using a full heuristic bootstrap analysis with 1000 replications. The genus Acicarpha (Calyceraceae) served as outgroup. RESULTS AND DISCUSSION Parsimony analysis of the multi-locus data matrix produced a single tree (Fig. 1) that placed Chucoa ilicifolia neither close to Chuquiraga nor Stifftia nor Gochnatia. Instead, Chucoa was found sister to Onoseris onoseroides (Mutisioideae-Onoserideae) with strong (100%) bootstrap support, highlighting Ferreyra s observation. The Onoseris clade, including the type species of Chucoa and Onoseris, is also statistically supported (100% bootstrap), warranting the inclusion of Chucoa in the genus Onoseris. To formalize this transfer, the following combination is proposed: Onoseris ilicifolia (Cabrera) Panero, comb. nov. Basionym Chucoa ilicifolia Cabrera, Bol. Soc. Argent. Bot. 6: Type: Peru: La Libertad, Prov. Santiago de Chuco, Angasmarca-Tulpo, A. López Miranda 1090 (holotype: LP). A second species of Chucoa, C. lanceolata, has been recently proposed (Sancho et al., 2005). The placement of this taxon awaits molecular studies. ACKNOWLEDGMENTS I thank M. Dillon of the Field Museum for material of Chucoa ilicifolia and Tom Wendt, TEX-LL herbarium curator, for locating specimens of Chucoa. I thank Bonnie S. Crozier and B. L. Turner for reviewing the manuscript and B. León and A. Cano for copies of R. Ferreyra's publications and photos of the holotype of C. lanceolata.

4 540 Phytologia (December 2009) 91(3) Sequencing of the matk, ndhd, ndhf, rbcl and rpob genes was funded by grant NSF DEB Fig.1. Single most parsimonious tree based on the MP analysis of five concatenated chloroplast genes. 422 of the 13,209 nucleotide sites were parsimony informative. Numbers above branches indicate percent bootstrap support values. LITERATURE CITED Cabrera, A. L Un nuevo género de Mutiseas del Perú. Bol. Soc. Argent. Bot. 6: Ferreyra, R Weberbaueriella, un nuevo género para las compuestas del Perú. Bol. Soc. Peruana Bot. 5: 1-6.

5 Phytologia (December 2009) 91(3) 541 Ferreyra, R Notas sobre especies peruanas de los géneros Arnaldoa, Chucoa (Compositae) y Monnina (Polygalaceae). 8: Freire, S. E., L. Katinas, and G. Sancho Gochnatia (Asteraceae: Mutisieae) and the Gochnatia complex: taxonomic implications from morphology. Ann. Missouri Bot. Gard. 89: Katinas, L., J. Pruski, G. Sancho and M. C. Tellería, The subfamily Mutisioideae (Asteraceae). Bot. Rev. 74: Sancho, G., S. E. Freire, L. Katinas and M. C. Tellería A new combination and a new species of Andean Mutisieae (Asteraceae). Taxon 54: Panero, J. L. and B. S. Crozier Primers for PCR amplification of Asteraceae chloroplast DNA. Lundellia 6: 1-9. Panero, J. L Shared molecular signatures support the inclusion of Catamixis in subfamily Pertyoideae (Asteraceae). Phytologia 90: Panero, J. L. and V. A. Funk The value of sampling anomalous taxa in phylogenetic studies: major clades of the Asteraceae revealed. Mol. Phylogenet. Evol. 47: Appendix 1. Voucher information and GenBank Accession numbers for sequences of Onoseris ilicifolia, Onoseris onoseroides, and Urmenetea. Voucher information listed in the following order: taxon name, collection, country of origin, herbarium. Genbank numbers listed in the following order: matk, ndhd, ndhf, rbcl, rpob. Onoseris ilicifolia (Cabrera) Panero, Sagástegui et al , Peru, F. GQ890332, GQ890335, GQ890338, GQ890341, GQ Onoseris onoseroides (H.B.K.) B. L. Rob., McVaugh 22341, Mexico, TEX. GQ890333, GQ890336, GQ890339, GQ890342, GQ Urmenetea atacamensis Phil., Panero & Crozier 8446, Chile, TEX. GQ890334, GQ890337, GQ890340, GQ890343, GQ